J. Nannoplankton Res. 31 (2), 2010, pp.58-112 © 2010 International Nannoplankton Association, Inc.
ISSN 1210-8049 Printed by The Sheridan Press, Hanover, PA, USA

58

A guide to extant coccolithophores (Calcihaptophycidae,
Haptophyta) using light microscopy
Miguel FradaEPPO (Evolution du Plancton & PaléoOcéans), Centre National de la Recherche Scientifique & Université Pierre & Marie Curie (UMR 7144),
Station Biologique, 29682 Roscoff, France and Centro de Geologia, Faculdade de Ciências, Universidade de Lisboa, Edifício C6, Campo Grande,
1749-016 Lisboa, Portugal; currently, Environmental Biophysics & Molecular Ecology Program, Institute of Marine & Coastal Sciences, Rutgers
University, 71 Dudley Road, New Brunswick, NJ 08901, USA; frada@marine.rutgers.edu

Jeremy Young
Palaeontology Department, The Natural History Museum, Cromwell Road, London, SW7 5BD, UK

Mário Cachão
Centro de Geologia, Faculdade de Ciências, Universidade de Lisboa, Edifício C6, Campo Grande, 1749-016 Lisboa, Portugal

Sílvia Lino, Ana Martins
Departamento de Oceanografia e Pescas (DOP), Universidade dos Açores, PortugalÁurea Narciso
Centro de Geologia, Faculdade de Ciências, Universidade de Lisboa, Edifício C6, Campo Grande, 1749-016 Lisboa, Portugal

Ian Probert, Colomban de Vargas
EPPO (Evolution du Plancton & PaléoOcéans), Centre National de la Recherche Scientifique & Université Pierre & Marie Curie (UMR 7144),
Station Biologique, 29682 Roscoff, France

Manuscript received 13th April, 2009; revised manuscript accepted 3rd June, 2010

Abstract We present here a collection of light microscope, and comparative scanning electron microscope, imagesof extant coccolithophores, sampled from various oceanic locations, and also from cultured strains. This series of images is intended to provide students and researchers interested in extant coccolithophore biology with an accessiblemeans of identifying the common coccolithophore species found in modern assemblages.

Keywords Coccolithophores, extant, diversity, light microscopy, scanning electronic microscopy
1. Intoduction
Coccolithophores include all algae from the Phylum Haptophyta that possess calcified scales (coccoliths) covering
the cell surface, at least in one phase of their life-cycle. According to the fossil record, the coccolithophores originatedin the Late Triassic (Bown, 1998), ~225Ma, since when
they have evolved and diversified, today comprising ~200
morphospecies (Jordan & Chamberlain, 1997; Young et al.,2003) within the monophyletic subclass Calcihaptophycidae (de Vargas et al., 2007; Liu et al., 2009).
Coccolithophores constitute a major component ofplanktonic communities throughout the world’s oceans(Okada & McIntyre, 1977). A single freshwater coccolithophore, Hymenomonas roseola, has also been documented (Manton & Peterfi, 1969). Coccolithophores are of
remarkable interest to a wide range of scientists. For marine biologists and oceanographers, they are among themain primary producers, and play a distinct role in theoceans’ ecosystems (Field et al., 1998; Balch, 2004). Forbiogeochemists, they are significant in the global carbon
and sulphur cycles (and therefore climate regulation)
through direct involvement in ocean-atmosphere gas exchange, and contribute to the transport of matter to theocean floor (Rost et al., 2003; Malin & Steinke, 2004). For
palaeontologists and geologists, the remarkably complete
and continuous coccolithophore fossil record (e.g. Bown
et al., 2004) makes them ideal tools for biostratigraphic,
evolutionary and palaeoceanographic studies of Mesozoic
and Cenozoic sediments.
Coccolithophores are characterised by a life-cycle composed of two phases that are morphologically distinct (heteromorphy) and have different ploidy levels, one being
haplontic and the other diplontic (haplo-diploidy; Billard,
1994; Billard & Inouye, 2004; Houdan et al., 2004). This
inference is based on limited data from chromosome counts
and ploidy-level comparisons made on coccolithophore
cultured strains (e.g. Gayral & Fresnel, 1983; Green et al.,
1996; Houdan et al., 2004) and supplemented through therecord of ‘combination coccospheres’ observed in natural
samples, these representing the transition between lifecycle phases, and bearing coccoliths characteristic of eachstage (e.g. Kleijne, 1991; Thomsen et al., 1991; Cros et al.,
2000; Geisen et al., 2002). From these studies, four typesof life-cycle associations have been identified in coccolithophores:
1. Diploid phase bearing heterococcoliths (coccoliths com-59

Frada et al.

posed of cycles of one or more radial arrays of elaborate
and variably-shaped crystal-units: Young et al., 1992;
Young et al., 2003) associated with a non-calcified haploidphase - Noelaerhabdaceae, Hymenomonadaceae and Pleurochrysidaceae;
2. Diploid phase bearing heterococcoliths associated witha haploid phase bearing holococcoliths (disc- or domeshaped coccoliths composed of numerous minute, equidimensional, rhombohedral calcite crystallites) Calcidiscaceae, Coccolithaceae, Helicosphaeraceae, Pontosphaeraceae, Syracosphaeraceae, Rhabdosphaeraceae
and Papposphaeraceae (Young et al., 2003; Frada et al.,2009);
3. One phase bearing heterococcoliths (possibly diploid)
associated with a phase bearing aragonitic coccoliths (possibly haploid) - Alisphaeraceae (Cros et al., 2000);
4. One phase bearing heterococcoliths (possibly diploid)
associated with a phase bearing nannoliths, being possibly
the haploid phase - Ceratolithaceae (Alcober & Jordan,
1997; Young et al., 2003).
Cross-polarised light microscopy is extensively used
for observing, identifying and describing coccoliths in fossil samples by palaeontologists (Bown, 1998). This powerful method has unfortunately been much less widelyadopted by biologists studying living coccolithophores, in
opposition to other light or electron microscopy techniques.
Calcite is highly birefringent and causes coccoliths to show
bright patterns (extinction figures), which allows them tobe readily differentiated from non-calcified micro-organisms. Moreover, as a result of their size, composition andstructure, different coccoliths generally show very distinct
extinction figures, which allow identification of many morphological species or genera, and also life-cycle phases(e.g. Bown, 1998; Young et al., 2004).
Here, we present a series of plates combining cross-polarised (XPL), differential-interference-contrast (DIC) andphase-contrast (PC) light microscope (LM) images of a variety of coccolithophore morphospecies (in some cases,
both life-cycle phases), collected from various oceanic locations and laboratory cultures. Scanning electron microscopy (SEM) images from the collection produced for
the Plankton*Net website (http://planktonnet.sb-roscoff.fr)
were added to each plate, in order to provide a cross-reference for recognition of each morphospecies. Our objective
is to provide the community interested in extant coccolithophore ecology with a guide for easy identification andtaxonomic assignment of coccolithophores from modern
samples. This publication complements the recent monographs of Cros & Fortuño (2002), Young et al. (2003) andMalinverno et al. (2008), by providing extensive LM documentation of extant coccolithophores. For detailed descriptions of coccolith morphology and taxonomy, theabove-mentioned monographs should be consulted.
2. Material and methods
The samples from which the images were taken were collected by several of the co-authors using a variety of meth-ods. The methodology is presented below according to thesample source.
2.1 Atlantic Meridional Transect (AMT)
16 (May-July, 2005), HOTS station,
Hawaii, central Pacific (June, 2005),
Tansei Maru KT11-06 cruise, Japaneseeast coast (May-June, 2006), bay ofVillefranche-sur-Mer, Mediterranean Sea(September, 2006), Belgica cruise, North
Atlantic (May-June, 2007)
Samples from these various locations were collected eitherwith a 5μm-mesh nannoplankton net from surface-waters
or, using Niskin bottles, from water at different depths (forfurther details on AMT16, see Robinson et al., 2006; Poulton et al., 2007). The samples, prepared for COD-FISH
morphogenetic analyses (Frada et al., 2006), were fixed for
1hr at 4˚C with 1% paraformaldehyde (pH8), filtered onto
0.2μm-pore-sized Whatman Anodisc membrane filters, anddried at room temperature. Later, a representative segmentof the filters was soaked in immersion oil (Olympus 04),
mounted between a glass slide and a coverslip and observed mainly in XPL, but for some species, PC or DIC
were also used (as indicated in the plate captions). Observations were performed with an Olympus BX51 microscope and the images were acquired with an RT-Slider Spotcooled charge coupled device digital camera.
2.2 Azores (May-July, 2008)
These samples were collected offshore of Horta, Faial island, Azores (38˚36’N, 28˚42’W) from surface-waters
using Niskin bottles. Samples were directly filtered onto
0.8μm-pore-sized Whatman cellulose nitrate membranes,and dried at room temperature. Slide preparation and observations were performed as described above.
2.3 Gulf of Naples (24th November, 2006)
These samples were collected with 5μm-mesh plankton
nets from surface-waters at an offshore station in the
Tyrrhenian Sea (39˚30’N, 13˚30’E). Samples were fixedwith formaldehyde at a final concentration of 2%. Observations were made with a Zeiss Axiovert microscope,equipped with a Zeiss Axiocam digital camera. For determination of the crystallographic orientation of the holococcolith crystals, an aliquot of the sample was filteredonto a 1μm-pore-sized Whatman cellulose acetate filter,permanently mounted using Norland optical adhesive(No.74), and examined under XPL using a Zeiss Axioplan
photomicroscope.
2.4 Sediment samples
Sediment samples were taken from the Saldanha hydrothermal field (Azores), from Core MD95-2040, recovered off the Portuguese margin (40˚34.91’N, 09˚51.67’W),
and from the beach in front of the Station Biologique,
Roscoff (France). For sample examination, smear-slides
were prepared, and observed under an optical polarisingGuide to extant coccolithophores

microscope (Olympus BX40 and BX51), at 1250x magnification. Digital images were taken with either an Olympus DP11 camera or an RT-Slider Spot cooled charge
coupled device camera. For further information on the sample collection and preparation see Parente et al. (2004) andNarciso et al. (2006).
3. The images
All scale-bars are 5μm, except where indicated on theplates. ‘NHM’ is the Natural History Museum, London;[HET] - heterococcolithophore, [HOL] - holococcolithophore. NB Copies of most of the SEM images are available
onthePlankton*Net
website(http://
planktonnet.sb-roscoff.fr). All taxonomic references can befound in Bown (1998), Young et al. (2003) and/or Malinverno et al. (2008).
3.1 Image index
Calcidiscaceae
Calcidiscus leptoporus (Murray & Blackman, 1898) Loeblich & Tappan, 1978 - Pl.8
Calcidiscus quadriperforatus (Kamptner, 1937) Quinn et al., 2004 [HET] (previously C. leptoporus large morphotype) - Pl.8
Calcidiscus quadriperforatus Quinn & Geisen in Saez et al., 2003 [HOL] (previously Syracolithus quadriperforatus Kamptner, 1937, combination establishedby Geisen et al., 2002) - Pl.9
Oolithotus fragilis (Lohmann, 1912) Martini & Müller, 1972 [HET] - Pl.10
Umbilicosphaera foliosa (Kamptner, 1963 ex Kleijne, 1993) Geisen in Saez et al.,2003 [HET] - Pl.12
Umbilicosphaera hulburtiana Gaarder, 1970 [HET] - Pl.12
Umbilicosphaera sibogae (Weber-van Bosse, 1901) Gaarder, 1970 [HET] - Pl.11
Calciosoleniaceae
Calciosolenia brasiliensis (Lohmann, 1902) Young et al., 2003 [HET] - Pl.35
Calciosolenia murrayi Gran, 1912 [HET] - Pl.34
Coccolithaceae
Coccolithus braarudii (Gaarder, 1962) Geisen et al., 2002 [HET] & [HOL] - Pls 4,
5
Coccolithus pelagicus (Wallich, 1877) Schiller, 1930 [HET] - Pl.6
Coccolithus pelagicus subsp. azorinus Parente & Cachão, 2004 in Parente et al.,2004 - Pl.7
Helicosphaeraceae
Helicosphaera carteri (Wallich, 1877) Kamptner, 1954 [HET] & [HOL] - Pls 15, 16
Helicosphaera hyalina Gaarder, 1970 [HET] - Pl.18
Helicosphaera wallichii (Lohmann, 1902) Okada & McIntyre, 1977 [HET] & [HOL]- Pl.17
Incertae sedis
Alisphaera ordinata (Kamptner, 1941) Heimdal, 1973 [HET] - Pl.40
Alisphaera sp. [HET] - Pl.40
Braarudosphaera bigelowii (Gran & Braarud, 1935) Deflandre, 1947 - Pl.41
Ceratolithus cristatus Kamptner, 1950 [HET] & [CER] - PlS 44, 45
Ceratolithus cristatus Kamptner, 1950 coccolithomorpha-type [HET] - Pls 44, 45,
46
Ceratolithus cristatus Kamptner, 1950 cristatus-type [CER] - Pl.45
Ceratolithus cristatus Kamptner, 1950 nishidae-type [HET] - Pls 44, 45
Florisphaera profunda Okada & Honjo, 1973 - Pl.43
Gladiolithus flabellatus (Halldal & Markali, 1955) Jordan & Chamberlain, 1993 Pl.47
Umbellosphaera irregularis Paasche in Markali & Paasche, 1955 [HET] - Pl.42
Umbellosphaera tenuis (Kamptner, 1937) Paasche in Markali & Paasche, 1955
[HET] - Pl.42
Umbellosphaera tenuis type IV (Kamptner, 1937) Paasche in Markali & Paasche,1955 - Pl.42
Noelaerhabdaceae
Emiliania huxleyi (Lohmann, 1902) Hay & Mohler in Hay et al., 1967 - Pl.1
Gephyrocapsa ericsonii McIntyre & Bé, 1967 - Pl.2
Gephyrocapsa muellerae Bréhéret, 1978 - Pl.2
Gephyrocapsa oceanica Kamptner, 1943 - Pl.2
Reticulofenestra sessilis (Lohmann, 1902) Jordan & Young, 1990 - Pl.3
Pleurochrysidaceae

60

Pleurochrysis carterae (Braarud & Fagerland, 1946) Christensen, 1978 var. carterae- Pl.13
Pleurochrysis placolithoides Fresnel & Billard, 1991 - Pl.14
Pontosphaeraceae
Pontosphaera japonica (Takayama, 1967) Nishida, 1971 [HET] & [HOL] - Pl.26
Pontosphaera multipora (Kamptner, 1948) Roth, 1970 [HET] - Pl.25
Pontosphaera syracusana Lohmann, 1902 - Pl.24
Scyphosphaera apsteinii Lohmann, 1902 [HET] & [HOL] - Pls 19-22
Scyphosphaera porosa Kamptner, 1967 [HET] - Pl.23
Rhabdosphaeraceae
Acanthoica quattrospina Lohmann, 1903 [HET] - Pl.38
Acanthoica sp. [HET] - Pl.38
Algirosphaera robusta (Lohmann, 1902) Norris, 1984 [HET] - Pl.39
Discosphaera tubifera (Murray & Blackman, 1898) Ostenfeld, 1900 [HET] - Pl.36
Palusphaera sp.1 Cros & Fortuño, 2002 [HET] - Pl.38
Rhabdosphaera clavigera Murray & Blackman, 1898 [HET] - Pl.37
Syracosphaeraceae
Coronosphaera binodata (Kamptner, 1927) Gaarder, 1977 [HET] - Pl.30
Coronosphaera mediterranea (Lohmann, 1902) Gaarder in Gaarder & Heimdal,
1977 [HET] - Pl.30
Michaelsarsia adriaticus (Schiller, 1930) Manton et al., 1984 [HET] - Pls 31, 32
Michaelsarsia elegans Gran, 1912 emend. Manton et al., 1984 [HET] - Pl.31
Ophiaster formosus (Gran, 1912) emend. Manton & Oates, 1983 [HET] - Pl.33
Ophiaster sp. [HET] - Pl.33
Syracosphaera anthos (Lohmann, 1912) Janin, 1987 [HET] & [HOL] - Pl.28
Syracosphaera pulchra Lohmann, 1902 [HET] - Pl.27
Syracosphaera sp. [HET] - Pl.29

Acknowledgements
This work was supported by a PhD Fellowship awarded to MF
by the Fundacão Para a Ciência e para a Tecnologia, Portugal, andan ATIP Grant awarded to CdV by the Centre National de laRecherche Scientifique, France. It is part of the multidisciplinary
project BOOM (Biodiversity of Open Ocean Microcalcifiers),
funded by the French Agence Nationale de la Recherche, Grant
ANR-05-BIODIV-004. We are grateful to numerous colleagues
for supplying samples and/or SEM images, including Lluïsa Cros,
Claire Findlay, Jacqueline Fresnel, Ric Jordan, Markus Geisen,
Annelies Kleijne, Vita Pariente, Isabella Percopo and Claudia
Sprengel. Constructive reviews by Lluïsa Cros Miguel and Ric
Jordan were very helpful to the final version of the manuscript
and are gratefully acknowledged.

References
Alcober, J. & Jordan, R.W. 1997. An interesting association between Neosphaera coccolithomorpha and Ceratolithus cristatus (Haptophyta). European Journal of Phycology, 32: 91-93.
Balch, W.M. 2004. Re-evaluation of the physiological ecology of
coccolithophores. In: H.R. Thierstein & J.R. Young (Eds).
Coccolithophores: From molecular processes to global impact. Springer Verlag: 165-190.
Billard, C. 1994. Life Cycles. In: J.G.B. Leadbeater (Ed.). The
Haptophyte Algae. Clarendon Press: 167-186.
Billard, C. & Inouye, I. 2004. What’s new in coccolithophore biology? In: H.R. Thierstein & J.R. Young (Eds). Coccolithophores: From molecular processes to global impact.
Springer Verlag: 1-30.
Bown, P. (Ed.) 1998. Calcareous nannofossil biostratigraphy.
Kluwer Academic Publishers: 315pp.
Bown, P.R., Lees, J.A. & Young, J.R. 2004. Calcareous nannoplankton evolution and diversity through time. In: H.R. Thierstein & J.R. Young (Eds). Coccolithophores: From molecular
processes to global impact. Springer Verlag: 481-508.
Couapel, M.J.J., Beaufort, L. & Young, J.R. 2009. A new Helicosphaera - Syracolithus combination coccosphere (Haptophyta) from the western Mediterranean Sea. Journal of

61

Frada et al.

Phycology, 45(4): 914-916.
Cros, L. & Fortuño, J.-M. 2002. Atlas of Northwestern Mediterranean Coccolithophores. Scientia Marina, 66: 1-186.
Cros, L., Kleijne, A., Zeltner, A., Billard, C. & Young, J.R. 2000.
New examples of holococcolith-heterococcolith combination
coccospheres and their implications for coccolithophorid biology. Marine Micropaleontology, 39(1-4): 1-34.
Field, C.B., Behrenfeld, M.J., Randerson, J.T. & Falkowski, P.
1998. Primary production of the biosphere: integrating terrestrial and oceanic components. Science, 281: 237-240.
Frada, M., Not, F., Probert, I. & de Vargas, C. 2006. CaCO3 optical detection with fluorescent in situ hybridization: a new
method to identify and quantify calcifying microorganisms
from the oceans. Journal of Phycology, 42(6): 1162-1169.
Frada, M., Percopo, I., Young, J., Zingone, A., de Vargas, C. &Probert, I. 2009. First observations of heterococcolithophoreholococcolithophore life cycle combinations in the Family
Pontosphaeraceae (Calcihaptophycideae, Haptophyta). Marine Micropaleontology, 71: 20-27.
Gayral, P. & Fresnel, J. 1983. Description, sexualité et cycle dedéveloppement d’une nouvelle Coccolithophoracée (Prymnesiophyceae): Pleurochrysis pseudoroscoffensis sp. nov.
Protistologica, 19: 245-261.
Geisen, M., Billard, C., Cros, L., Probert, I. & Young, J. 2002.
Life cycle associations involving pairs of holococcolithophorid species: Intraspecific variation or cryptic speciation? European Journal of Phycology, 37: 531-550.
Green, J.C., Course, P.A. & Tarran, G.A. 1996. The life-cycle ofEmiliania huxleyi: A brief review and a study of relative
ploidy levels analysed by flow cytometry. Journal of Marine
Systems, 9(1-2): 33-44.
Houdan, A., Billard, C., Marie, D., Not, F., Saez, A., Young, J.R.
& Probert, I. 2004. Holococcolithophore-heterococcolithophore (Haptophyta) life cycles: flow cytometry analysis ofrelative ploidy levels. Systematics & Biodiversity, 1: 453-465.
Jordan, R.W. & Chamberlain A.H.L.1997. Biodiversity among
haptophyte algae. Biodiversity and Conservation, 6: 131-152.
Kleijne, A. 1991. Holococcolithophorids from the Indian Ocean,
Red Sea, Mediterranean Sea and North Atlantic Ocean. Marine Micropaleontology, 17: 1-76.
Liu, H., Aris-Brosou, S., Probert, I. & de Vargas, C. 2009. A timeline of the environmental genetics of the haptophytes. Molecular Biology and Evolution, 27: 161-176.
Malin, G. & Steinke, M. 2004. Dimethyl Sulfide Production:
What is the Contribution of the Coccolithophores? In: H.R.
Thierstein & J.R. Young (Eds). Coccolithophores: From molecular processes to global impact. Springer Verlag: 127-164.
Malinverno, E., Dimiza, M.D., Triantaphyllou, M., Dermitzakis,
M. & Corselli, C. 2008. Coccolithophores of the Eastern
Mediterranen Sea: A look into the marine microworld. Ion
Publishing Group, Peristeri: 188pp.
Manton, I. & Peterfi, L.S. 1969. Observations on the fine structure of coccoliths, scales and the protoplast of a freshwatercoccolithophorid, Hymenomonas roseola Stein, with supplementary observations on the protoplast of Cricosphaera
carterae. Proceedings, Royal Society, London, 172: 1-15.
Narciso, A., Cachão, M & Abreu, L. 2006. Coccolithus pelagicus
subsp. pelagicus versus Coccolithus pelagicus subsp.
braarudii (Coccolithophore, Haptophyta): A proxy for surface subarctic Atlantic waters off Iberia during the last 200kyr.
Marine Micropaleontology, 59: 15-34.
Okada, H. & McIntyre, A. 1977. Modern coccolithophores of the
Pacific and North Atlantic Oceans. Micropaleontology, 23(1):1-55.
Parente, Á., Cachão, M., Baumann, K.-H., Abreu, L. & Ferreira,J. 2004. Morphometry of Coccolithus pelagicus s.l. (Coccolithophore, Haptophyta) from offshore Portugal, during the last
200kyr. Marine Micropaleontology, 50(1): 107-120.
Poulton, A.J., Adey, T.R., Balch, W.M. & Holligan, P.M. 2007.
Relating coccolithophore calcification rates to phytoplankton
community dynamics: Regional differences and implications
for carbon export. Deep-Sea Research II, 54(5-7): 538-557.
Robinson, C., Poulton, A.J., Holligan, P.M., Baker, A.R., Forster,G., Gist, N., Jickells, T.D., Malin, G., Upstill-Goddard, R.,
Williams, R.G., Woodward, E.M.S. & Zubkov, M.V. 2006.
The Atlantic Meridional Transect (AMT) Programme: A contextual view 1995-2005. Deep-Sea Research II, 53(14-16):1485-1515.
Rost, B., Riebesell, U., Burkhardt, S. & Sultemeyer, D. 2003. Carbon acquisition of bloom-forming marine phytoplankton.
Limnology and Oceanography, 48(1): 55-67.
Thomsen, H.A., Østergaard, J.B., & Hansen, L.E. 1991. Heteromorphic life histories in Arctic coccolithophorids (Prymnesiophyceae). Journal of Phycology, 27: 634-642.
de Vargas, C., Aubry, M.-P., Probert, I. & Young J.R. 2007. Origin and Evolution of Coccolithophores: From Coastal Hunters
to Oceanic Farmers. In: P. Falkowski & A.H. Knoll (Eds).
Evolution of Aquatic Photoautotrophs. Elsevier: 251-286.
Young, J.R. 2008. Scyphosphaera porosa Kamptner, 1967 rediscovered in the plankton. J. Nannoplankt. Res., 30(1): 35-38.
Young, J.R., Didymus, J.M., Bown, P.R., Prins, B. & Mann, S.
1992. Crystal assembly and phylogenetic evolution in heterococcoliths. Nature, 356: 516-518.
Young, J.R., Geisen, M., Cros, L., Kleijne, A., Sprengel, C.,
Probert, I. & Østergaard, J. 2003. A guide to extant coccolithophore taxonomy. J. Nannoplankt. Res., Spec. Iss, 1:
125pp.
Young, J.R., Henriksen, K. & Probert, I. 2004. Structure and morphogenesis of the coccoliths of the CODENET species. In:
H.R. Thierstein & J.R. Young (Eds). Coccolithophores: From
molecular processes to global impact. Springer Verlag: 191216.

Plate 1
1-5, 7-10. Emiliania huxleyi diploid phase [HET]1. 50m water-depth; Canary Islands, North Atlantic, 29˚41’N,
17˚53’W; SEM (image NHM 118-14)
2. Surface waters; South Atlantic, 04˚S, 25˚W; AMT16 cruise,
June, 2005; XP
3. 30m water-depth; Faial, Azores, 30˚32’N, 28˚33’W; May,
2008; XP
4. Surface waters; Japan, 40˚N, 145˚E; Tansei Maru KT11-06
cruise, May, 2006; XP
5. Culture RCC 1249; DIC
7. Detached coccolith; surface waters; South Atlantic, 04˚S,
25˚W; AMT16 cruise, June 2005; XP
8-10. Series of images from different phases of an E. huxleyi
diploid-phase bloom, English Channel, Belgica cruise, May, 2007
8. Exponentially-growing phase - healthy cells and a few detachedcoccoliths; XP
9. Stationary phase - increased abundance of detached coccoliths,
appearance of clumps of cellular debris; XP
10. End of the bloom - cellular debris and detached liths, very few
whole coccospheres; XP
6. E. huxleyi haploid biflagellated (arrowed) phase
Culture RCC 1249; DIC

Guide to extant coccolithophores

Plate 1
Noelaerhabdaceae

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Frada et al.

Plate 2
1-7. Gephyrocapsa oceanica diploid phase [HET]1. 5m water-depth; Canary Islands, North Atlantic, 27˚33’N, 13˚38’W; SEM (image NHM 104-8)
2-4. 30m water-depth; Faial, Azores, 30˚32’N, 28˚33’W; May, 2008; XP
5. Surface waters; Japan, 34˚26’N, 139˚E; Tansei Maru cruise, May, 2006; XP
6, 7. Detached coccoliths; Faial, Azores, 30˚32’N, 28˚33’W; May, 2008; XP
8-12. G. muellerae diploid phase [HET]8. 37m water-depth; Alboran Sea, western Mediterranean, 37˚23’N, 0˚56’W; SEM (image NHM 117-19)
9, 10, 12. 30m water-depth; Faial, Azores, 30˚32’N, 28˚33’W; May, 2008; XP
11. Detached coccolith; Faial, Azores, 30˚32’N, 28˚33’W; June, 2008; XP
13, 14. G. ericsonii diploid phase [HET]13. 130m water-depth; HOTS station, Hawaii, Pacific Ocean, 22˚45’N, 158˚E; SEM (image NHM 217-17)
14. Surface water; Faial, Azores, 30˚32’N, 28˚33’W; June, 2008; XP

Guide to extant coccolithophores

Plate 2
Noelaerhabdaceae

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Frada et al.

Plate 3
Noelaerhabdaceae

Plate 3
1-5. Reticulofenestra sessilis [HET]
1. Cells associated with centric diatom Thalassiosira sp.; 175m water-depth; Gulf of Mexico, 26˚19’N, 59˚38’W; SEM (image NHM
139-47)2-5. Cells associated with unidentified diatom; surface waters; South Atlantic, 31˚49’S, 01˚30’W; AMT16 cruise, May, 2005; XP (2,4), PC (3, 5)

Plate 41, 3-5. Coccolithus braarudii [HET]1. 3200m water-depth; North Atlantic, 48˚N, 20˚W; SEM (image NHM 185-24)
3-5. Culture strain RCC1197; XP
2, 6, 7. C. braarudii [HOL] ‘Crystallolithus braarudii’ haploid phase
2. Collapsed coccosphere; Alboran Sea, western Mediterranean; SEM (image from L. Cros, CSIC-ICM, Barcelona)
6, 7. Culture strain RCC1197; XP

Guide to extant coccolithophores

Plate 4
Coccolithaceae

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Plate 5
Coccolithaceae

1-6. Coccolithus braarudii [HET]1, 2. Detached coccoliths; sediment sample, Roscoff, France; PC(1), XP (2)3. Early stage of coccolith formation; sediment sample, Roscoff,France; XP
4-6. Culture RCC1197, dissociated coccospheres; XPGuide to extant coccolithophores

Plate 6
Coccolithaceae

1, 2. Coccolithus pelagicus [HET]1. Surface waters; Iceland, 63˚27’N, 20˚12’W; SEM (image NHM 111-30)
2. 30m water-depth; Faial, Azores, 30˚32’N, 28˚33’W; May, 2008; XP

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Plate 7
Coccolithaceae

1-5. Coccolithus pelagicus subsp. azorinus
1-4. Detached coccoliths, distal view; sediment sample; PC (a), XP (b)5. Coccosphere; sediment sample; PCGuide to extant coccolithophores

Plate 8
Calcidiscaceae

1-3. Calcidiscus quadriperforatus [HET] (previously C. leptoporus large morphotype)
1. 20m water-depth; eastern equatorial Atlantic; 18˚N, 17˚W; SEM (image NHM 240-36)
2. Surface waters; South Atlantic, 01˚37’S, 24˚59’W; AMT16 cruise, June, 2005; XP
3. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP
4-7. C. leptoporus [HET] intermediate morphotype
4. 5m water-depth; Canary Islands, North Atlantic, 33˚27’N, 9˚10’W; SEM (image NHM 103-74)
5-7. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, May, 2005; XP
8, 9. C. leptoporus [HET] small morphotype
8. Off Namibia, South Atlantic; SEM (image NHM 136-03)
9. Possible small morphotype; 30m water-depth; Faial, Azores, 30˚32’N, 28˚33’W; July, 2008; DIC

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Plate 9Calcidiscaceae

Plate 91-4. Calcidiscus quadriperforatus [HOL]1. 25m water-depth; Canary Islands, North Atlantic, 29˚45’N, 17˚56’W; SEM (image NHM 113-17)
2-4. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP (2), DIC (3, 4)

Plate 10
1-5. Oolithotus fragilis [HET]1. 37m water-depth; Alboran Sea, western Mediterranean, 37˚23’N, 0˚56’W; SEM (image NHM 119-76)
2, 4. Culture strain RCC 1482; XP
3. Dispersed coccoliths; XP
5. Group of four cells; culture strain RCC 1482; XPGuide to extant coccolithophores

Plate 10
Calcidiscaceae

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Plate 11
Calcidiscaceae

1-6. Umbilicosphaera sibogae [HET]1. 29m water-depth; eastern equatorial Atlantic, 18˚00’N, 26˚59’W; SEM (image NHM 240-09)
2-5. Surface waters; AMT16 cruise, June, 2005; XP
2. South Atlantic, 01˚37’S, 24˚59’W
3. South Atlantic, 20˚11’S, 24˚59’W
4. North Atlantic, 04˚16’N, 27˚01’W
5. Detached coccolith, South Atlantic; 20˚11’S, 24˚59’W
6. 30m water-depth; Faial, Azores, 30˚32’N, 28˚33’W; May, 2008; XP

Guide to extant coccolithophores

Plate 12
Calcidiscaceae

1-3. Umbilicosphaera foliosa [HET]1. Surface waters; Ibo Port, Miyake Island, Japan, 34˚06’N, 139˚30’E; SEM (image NHM 129-10)
2. Surface waters; South Atlantic, 26˚49’S, 10˚30’W; AMT16 cruise, May, 2005; XP
3. Surface waters; North Atlantic, 18˚57’N, 34˚12’W; AMT16 cruise, May, 2005; XP
4-6. U. hulburtiana [HET]4. 5m water-depth; North Atlantic, 21.91˚N, -20.29˚E; SEM (image NHM 114-34)
5. Surface waters; South Atlantic, 22˚52’S, 24˚59’W; AMT16 cruise, May, 2005; XP
6. 70m water-depth; South Atlantic, 01˚37’S, 24˚59’W; AMT16 cruise, May, 2005; XP

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Plate 13
Pleurochrysidaceae

1-3. Pleurochrysis carterae var. carterae
1. Collapsed coccosphere; SEM (image Fr 99238 from J. Fresnel, University of Caen, France)
2, 3. Cultured strain RCC 1402; XPGuide to extant coccolithophores

Plate 14
Pleurochrysidaceae

1-4. Pleurochrysis placolithoides
1. Collapsed coccosphere; culture strain RCC 1401; SEM (image NHM 85-15)
2-4. Cultured strain AC59; XP

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Plate 15
1, 4. Helicosphaera carteri [HET]1. 5m water-depth; off Namibia, South Atlantic, 20˚S, 09˚E; SEM (image NHM 136-37)
4. Surface waters; North Atlantic, 10˚00’N, 29˚47’W; AMT16 cruise, June, 2005; XP
2, 5. H. carteri [HOL solid] (formerly Syracolithus catilliferus)
2. 50m water-depth; Canary Islands, North Atlantic, 29˚41’N, 17˚53’W; SEM (image P233B314 from Claudia Sprengel, Universityof Bremen)
5. Surface waters; North Atlantic, 38˚18’N, 30˚03’W; AMT16 cruise, June, 2005; XP
3, 7. H. carteri [HOL perforate] (formerly Syracolithus confusus)
3. 20m water-depth; Villefranche-sur-Mer, Mediterranean; SEM (image NHM 284-24)
7. Surface waters; North Atlantic, 38˚18’N, 30˚03’W; AMT16 cruise, June, 2005; XP
6. H. carteri [HET & HOL solid]
Combination coccosphere bearing both heterococcoliths and holococcoliths; surface waters; North Atlantic, 38˚18’N, 30˚03’W; AMT16
cruise, June, 2005; XP

Guide to extant coccolithophores

Plate 15
Helicosphaeraceae

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Plate 16
1, 6, 7. Helicosphaera carteri [HOL solid]
1. Distal and side views; North Atlantic, 38˚18’N, 30˚03’W; AMT16 cruise, June, 2005; XP
6. Surface waters; South Atlantic, 20˚93’S, 24˚59’W; AMT16 cruise, May, 2005; XP
7. Surface waters; North Atlantic, 38˚18’N, 30˚03’W; AMT16 cruise, June, 2005; XP
2, 3, 5. H. carteri [HET]2, 3. Detached coccoliths; surface waters; North Atlantic, 10˚N, 29˚47’W; AMT16 cruise, June, 2005; XP
5. Surface waters; North Atlantic, 10˚N, 29˚47’W; AMT16 cruise, June, 2005; XP
4, 8. H. carteri [HOL perforate]
4. Close-up of coccoliths (see fig.8); North Atlantic, 31˚22’N, 42˚08’W; AMT16 cruise, June, 2005; XP
8. Surface waters; North Atlantic, 31˚22’N, 42˚08’W; AMT16 cruise, June, 2005; XP
9. H. carteri [HOL solid & HOL perforate]
Combination coccosphere bearing two types of holococcolith; surface waters; North Atlantic, 38˚18’N, 30˚03’W; AMT16 cruise,
June, 2005; XP

Guide to extant coccolithophores

Plate 16
Helicosphaeraceae

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Plate 17
Helicosphaeraceae

1, 3-5. Helicosphaera wallichii [HET]1. Surface waters; Miyake Port, Miyake Island, Japan, 34˚01’N, 139˚30’E; SEM (image NHM116-74)
3-5. Surface waters; North Atlantic, 04˚16’N, 27˚01’W; AMT16 cruise, May, 2005; XP
2, 6, 7. H. wallichii [HOL] (formerly Syracolithus ponticuliferus, combination established by Couapel et al., 2009)
2. 10m water-depth; Gulf of Mexico; SEM (image NHM-CSF0112a)6, 7. Surface waters; North Atlantic, 25˚40’N, 37˚40’W; AMT16 cruise, May, 2005; XP

Guide to extant coccolithophores

Plate 18
Helicosphaeraceae

1-4. Helicosphaera hyalina [HET]
1. Culture specimen; SEM (image NHM187-51)
2, 3. Surface waters; Faial, Azores, 30˚32’N, 28˚33’W; XP (2), DIC (3)4. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP

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Plate 19
Pontosphaeraceae

1μm

1. Scyphosphaera apsteinii [HET]80m water-depth; HOTS station, Hawaii, South Pacific, 22˚45’N, 158˚E; SEM (image NHM 217-83)
2, 3. S. apsteinii [HOL] (formerly Syracolithus schillerii, combination established by Frada et al., 2009)
2. 80m water-depth; HOTS station, Hawaii, South Pacific, 22˚45’N, 158˚E; SEM (image NHM 217-77)
3. Close-up of coccoliths; surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise; SEM (image NHM 271-06)

Guide to extant coccolithophores

Plate 20
Pontosphaeraceae

1, 3, 4. Scyphosphaera apsteinii [HET]1. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; May, 2005; XP
3, 4. Detached coccoliths, distal view; North Atlantic, 36˚27’N, 36˚55’W; AMT16 cruise, June, 2005; XP
2, 5, 6. S. apsteinii [HOL]
2. Surface waters; North Atlantic, 15˚45’N, 32˚35’W; AMT16 cruise, May, 2005; XP
5-6. Detached holococcoliths, side and distal views; North Atlantic, 15˚45’N, 32˚35’W; AMT16 cruise, May, 2005; XP

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Plate 21
Pontosphaeraceae

1-4. Scyphosphaera apsteinii [HET & HOL]
Combination coccosphere; 140m water-depth; North Atlantic, 38˚18’N, 30˚03’W; AMT16 cruise, May, 2005; XP (1), DIC (2), PC (3),
SEM (4)5, 6. S. apsteinii [HET]5. Surface waters; North Atlantic, 36˚27’N, 36˚55’W; AMT16 cruise, June, 2005; XP
6. Surface waters; South Atlantic, 36˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP

Guide to extant coccolithophores

Plate 22
Pontosphaeraceae

1-6. Scyphosphaera apsteinii [HOL]
1, 4. Different focal planes, surface waters; North Atlantic, 15˚45’N, 32˚35’W; AMT16 cruise, June, 2005; XP
2, 3, 5, 6. Surface waters; AMT16 cruise, May, 2005
2. North Atlantic, 29˚09’N, 39˚32’W; XP
3, 5, 6. South Atlantic, 36˚31’S, 17˚13’W; XP (3), DIC (5, 6)

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Plate 23
Pontosphaeraceae

31-3. Scyphosphaera porosa [HET]1, 2. 96m water-depth; South Atlantic, 31˚49’S, 10˚30’E; AMT16
cruise, May, 2005; XP (1), DIC (2)3. Line-drawings of holotype (modified from Young, 2008)

Guide to extant coccolithophores

Plate 24
Pontosphaeraceae

1-4. Pontosphaera syracusana
1. 5m water-depth; Alboran Sea, western Mediterranean, 37˚27’N, 01˚28’W; SEM (image NHM 170-16)
2, 4. Surface waters; South Atlantic, 36˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP
3. Detached heterococcoliths; surface waters; South Atlantic, 01˚57’S, 24˚59’W; AMT16 cruise, June, 2005; XP

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Plate 25
Pontosphaeraceae

1-7. Pontosphaera multipora [HET]1. Coccolith, distal view; 120m water-depth; Gulf of Mexico, 26˚39’N, 39˚50’E; SEM (image NHM 134-06)
2. Coccoliths, distal view; 60m water-depth; Gulf of Mexico, 26˚19’N, 59˚38’W; SEM (image NHM CSF0103)
3. Surface waters; South Atlantic, 27˚49’S, 10˚30’W; AMT16 cruise, May, 2005; XP
4. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP
5-7. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, May, 2005; XP

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Plate 26
Pontosphaeraceae

1, 3. Pontosphaera japonica [HET]1. 80m water-depth; HOTS station, Hawaii, North Pacific, 22˚45’N, 158˚E; SEM (image NHM 217-82)
3. Surface waters; South Atlantic, 01˚37’S, 24˚59’W; AMT16 cruise, June, 2005; XP
2, 5. P. japonica [HET & HOL] (‘Syracolithus’-like holococcolithophore combination coccosphere, see Frada et al. (2009) for discussion)
Gulf of Naples; SEM (2, image from Isabella Percopo, SZN Naples), XP (5, image from Jeremy Young, NHM)
4, 6, 7. P. japonica [HOL]4, 6. Surface waters; eastern Tyrrhenian Sea, 39˚30’N, 13˚30’E; November, 2006; DIC (4), XP (6) (image from Jeremy Young, NHM)
7. Collapsed coccosphere; surface waters; North Atlantic, 18˚57’N, 34˚12’W; AMT16 cruise, May, 2005; XP

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Plate 27
Syracosphaeraceae

1-7. Syracosphaera pulchra [HET]1. 50m water-depth; Canary Islands, North Atlantic, 29˚41’N, 17˚53’E; SEM (image NHM 166-05)
2, 5. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June, 2005; XP
3. Surface waters; North Atlantic, 04˚16’N, 27˚01’W; AMT16 cruise, June, 2005; XP
4. Surface waters; South Atlantic, 22˚52’S, 24˚59’W; AMT16 cruise, May, 2005; XP
6, 7. Surface waters; Faial, Azores, 30˚32’N, 28˚33’W; May, 2008; XP

Guide to extant coccolithophores

Plate 28
Syracosphaeraceae

1, 3-5. Syracosphaera anthos [HET]1. 42.5m water-depth; Alboran Sea, western Mediterranean, 37˚25’N, 00˚25’W; SEM (image NHM 144-12)
3-5. Surface waters; South Atlantic, 31˚49’S, 01˚30’E; AMT16 cruise, May, 2005; XP (3, 4), DIC (5)2, 6, 7. S. anthos [HOL] (formerly Periphyllophora mirabilis, combination established by Cros et al., 2000)
2. 5m water-depth; Alboran Sea, western Mediterranean, 35˚54’N, 01˚31’W; SEM (image NHM 133-38)
6. Surface waters; Faial, Azores, 30˚32’N, 28˚33’W; AMT16 cruise, May, 2005; XP
7. Surface waters; South Atlantic, 26˚03’S, 17˚13’W; AMT16 cruise, May, 2005; XP

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Plate 29
Syracosphaeraceae

Plate 29
1-4. Syracosphaera sp. [HET]Surface waters; South Atlantic, 31˚49’S, 01˚30’E; AMT16 cruise, May, 2005; XP

Plate 30
1, 3-6. Coronosphaera mediterranea [HET]1. 5m water-depth; North Atlantic, 21˚16’N, 20˚41’W; SEM (image NHM 114-62)
3. Surface waters; North Atlantic, 25˚40’N, 37˚40’W; AMT16 cruise, June, 2005; XP
4, 5. Surface waters; Mediterranean, Villefranche-sur-Mer, France; March, 2007; XP
6. Surface waters; Faial, Azores, 30˚32’N, 28˚33’W; June, 2008; XP
2, 7, 8. C. binodata [HET]2. 5m water-depth; Canary Islands, North Atlantic, 33˚27’N, 09˚10’W; SEM (image NHM 103-33)
7. Detached coccoliths, distal view; Canary Islands, North Atlantic, 33˚27’N, 09˚10’W; SEM (image NHM 103-33)
8. Surface waters; South Atlantic, 22˚52’S, 24˚59’W; AMT16 cruise, June, 2005; XP

Guide to extant coccolithophores

Plate 30
Syracosphaeraceae

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Plate 31
Syracosphaeraceae

1, 3. Michaelsarsia elegans [HET]1. 42.5m water-depth; Alboran Sea, western Mediterranean, 37˚25’N, 00˚25’W; SEM (image NHM 127-23)
3. 50m water-depth; South Atlantic, 31˚04’S, 10˚30’E; AMT16 cruise, May, 2005; XP
2. M. adriaticus [HET]Surface waters; Miyake Island, Japan, 34˚06’N, 139˚30’E; SEM (image NHM 119-61)

Guide to extant coccolithophores

Plate 32
Syracosphaeraceae

1-3. Michaelsarsia adriaticus [HET]1, 3. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP
2. 10m water-depth; Japan, 34˚04’N, 140˚02’E; Tansei Maru cruise, May, 2006; XP

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Plate 33
Syracosphaeraceae

1. Ophiaster formosus [HET]42.5m water-depth; Alboran Sea, western Mediterranean, 37˚25’N, 00˚25’W; SEM (image NHM 125-14)
2-4. Ophiaster sp. [HET]2, 3. 96m water-depth; South Atlantic, 31˚49’S, 16˚28’E; AMT16 cruise, May, 2005; XP
4. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June, 2005; XP

Guide to extant coccolithophores

Plate 34
Calciosoleniaceae

1-4. Calciosolenia murrayi [HET]1. 37m water-depth; Alboran Sea, western Mediterranean, 37˚23’N, 00˚56’W; SEM (image NHM 117-29)
2-4. Surface waters; South & North Atlantic, 26˚31’S, 17˚13’W & 38˚18’N, 30˚03’W; AMT16 cruise, May & June, 2005; XP

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Plate 35
Calciosoleniaceae

1-5. Calciosolenia brasiliensis [HET]1. 5m water-depth; Alboran Sea, western Mediterranean, 35˚54’N, 01˚21’E; SEM (image NHM 189-31)
2, 4. 96m water-depth; South Atlantic, 31˚49’S, 16˚28’E; AMT16 cruise, May, 2005; XP
3. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP
5. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June, 2005; XP

Guide to extant coccolithophores

Plate 36
Rhabdosphaeraceae

1-5. Discosphaera tubifera [HET]1. 5m water-depth; Alboran Sea, western Mediterranean, 37˚25’N, 00˚25’E; SEM (image NHM 123-53)
2. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP
3, 4. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June 2005; XP
5. Surface waters; North Atlantic, 25˚40’N, 37˚40’W; AMT16 cruise, June, 2005; XP

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Plate 37
Rhabdosphaeraceae

1-5. Rhabdosphaera clavigera [HET]1. 5m water-depth; Alboran Sea, western Mediterranean, 37˚25’N, 00˚25’E; SEM (image NHM 124-02)
2, 3. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June, 2005; XP
4, 5. Surface waters; South Atlantic, 26˚31’S, 17˚13’W; AMT16 cruise, May, 2005; XP

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Plate 38
Rhabdosphaeraceae

1. Acanthoica quattrospina [HET]27m water-depth; Portuguese Shelf, North Atlantic, 38˚44’N, 09˚46’W; SEM (image NHM 121-22)
2. Palusphaera sp.1 [HET]2. Collapsed coccosphere; 87m water-depth; North Atlantic, 34˚17’N, 34˚23’W; SEM (image AK81-39 from Annelies Kleijne, VU
Amsterdam)3, 4. Palusphaera sp. [HET]Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June, 2005; XP
5, 6. Acanthoica sp. [HET]5. Surface waters; South Atlantic, 22˚52’S, 24˚59’W; AMT16 cruise, May, 2005; XP
6. 67m water-depth; North Atlantic, 36˚27’N, 36˚55’W; AMT16 cruise, May, 2005; XP103

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Plate 39
Rhabdosphaeraceae

1-6. Algirosphaera robusta [HET]1. Gulf of Mexico; SEM (image from Vita Pariente, Texas A&M University)2, 4, 5. Surface waters; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June, 2005; XP
3, 6. Surface waters; South Atlantic, 01˚37’S, 24˚59’W; AMT16 cruise, June, 2005; XP

Guide to extant coccolithophores

Plate 40
Incertae sedis

1. Alisphaera ordinata
5m water-depth; off Namibia, South Atlantic, 20˚52’S, 06˚15’E; SEM (image NHM 136-09)
2-4. Alisphaera sp. [HET]
30m water-depth; Faial, Azores, 30˚32’N, 28˚33’W; May, 2008; XP (2, 4), DIC (3)

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Plate 41
Incertae sedis

1-6. Braarudosphaera bigelowii
1. Surface waters; NE Atlantic, 41˚53’N, 09˚56’W; SEM (image NHM 209-28)
2. Collapsed coccosphere; Faial, Azores, 30˚32’N, 28˚33’W; June, 2008; XP
3, 4, 6. Surface waters; Faial, Azores, 30˚32’N, 28˚33’W; July, 2008; XP (3, 4 left, 6), DIC (4 right)
5. Detached nannolith; Faial, Azores, 30˚32’N, 28˚33’W; July, 2008; XP

Guide to extant coccolithophores

Plate 42
Incertae sedis

1. Umbellosphaera tenuis type IV
5m water-depth; Alboran Sea, western Mediterranean, 37˚25’N, 00˚25’W; SEM (image NHM 140-11)
2, 6, 7. U. irregularis [HET]2. 50m water-depth; HOTS station, Hawaii, North Pacific; 22˚45’N, 158˚E; SEM (image NHM 217-12)
6, 7. Surface waters; South Atlantic, 22˚27’S, 24˚59’W (6), 20˚11’S, 24˚59’W (7); AMT16 cruise, June, 2005; XP
3-5. U. tenuis [HET]39m water-depth; North Atlantic, 35˚05’N, 41˚51’W; AMT16 cruise, June, 2005; XP

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Plate 43
Incertae sedis

1-6. Florisphaera profunda
1. 175m water-depth; Gulf of Mexico, 26˚19’N, 59˚38’W; SEM (image NHM 132-08)
2. 50m water-depth; South Atlantic, 31˚58’S, 16˚58’E; AMT16 cruise, June, 2005; XP
3, 4. Surface waters, night; South Atlantic, 20˚11’S, 24˚59’W; AMT16 cruise, June, 2005; XP
5, 6. Surface waters, night; South Atlantic, 22˚52’S, 24˚59’W; AMT16 cruise, June, 2005; XP

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Plate 44
Incertae sedis

1. Ceratolithus cristatus [HET]50m water-depth; South Atlantic, 31˚04’S, 16˚28’E; AMT16 cruise, May, 2005; XP
2. C. cristatus nishidae type [HET]50m water-depth; South Atlantic, 31˚04’S, 16˚28’E; AMT16 cruise, May, 2005; XP
3-6. C. cristatus coccolithomorpha type [HET]3. Collapsed coccosphere?; 10m water-depth; South Atlantic, 31˚58’S, 16˚58’E; AMT16 cruise, May, 2005; XP
4-6. Same coccosphere viewed in different focal planes; 50m water-depth; South Atlantic, 31˚04’S, 16˚28’E; AMT16 cruise, May, 2005;
XP

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Plate 45
1. Ceratolithus cristatus cristatus type [CER] (according to Young et al., 2003, it is likely that the ceratolithid stage - CER - is equivalent to the holococcolith stage in other taxa, and is probably thus haploid)
Detached nannolith, proximal view; 10m water-depth; Gulf of Mexico, 50˚N, 04˚01’W; SEM (image NHM 132-3)
2, 5-9. C. cristatus [CER]
2. Detached nannolith, distal view; surface waters; Ibo Port, Miyake Island, Japan, 34˚06’N, 139˚30’E; SEM (image NHM 129-07)
5. Note presence of two horse-shoe-shaped nannoliths (arrow A) positioned more or less parallel to each other, but on opposite sides,
inside a common membrane-like structure (arrow B); 10m water-depth; South Atlantic, 31˚49’S, 10˚30’E; AMT16 cruise, June, 2005;
XP
6. 50m water-depth; South Atlantic, 31˚04’S, 16˚28’E; AMT16 cruise, May, 2005; XP
7, 8. Surface waters; South Atlantic, 31˚34’S, 09˚19’E; AMT16 cruise, May, 2005; XP
9. 50m water-depth; South Atlantic, 31˚58’S, 16˚58’E; AMT16 cruise, June, 2005; XP
3. C. cristatus coccolithomorpha type [HET] (= Neosphaera coccolithomorpha Lecal-Schlauder, 1950)
60m water-depth; Gulf of Mexico, 27˚36’N, 95˚E; SEM (image NHM CSF0146)
4. C. cristatus nishidae type [HET] (= Neosphaera coccolithomorpha var. nishidae Kleijne, 1993)
50m water-depth; Canary Islands, North Atlantic, 29˚41’N, 17˚53’W; SEM (image NHM P233B315)

Guide to extant coccolithophores

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Plate 46
Incertae sedis

1-4. Ceratolithus cristatus coccolithomorpha type [HET]1-3. 50m water-depth; South Atlantic, 31˚04’S, 16˚28’E; AMT16 cruise, May, 2005; XP (2, 3 taken in different focal planes)
4. 105m water-depth; South Atlantic, 25˚96’S, 21˚56’W; AMT16 cruise, May 2005; XP

Guide to extant coccolithophores

Plate 47
Incertae sedis

1-4. Gladiolithus flabellatus
1. 180m water-depth; HOTS station, Hawaii, North Pacific, 22˚45’N, 158˚E; SEM (image NHM 219-25a)2. 125m water-depth; Mediterranean Sea, Villefranche sur Mer; XP (Plankton*Net, ©Jeremy Young, NHM)
3. 96m water-depth; South Atlantic, 31˚49’S, 10˚30’E; AMT16 cruise, June, 2005; XP
4. 50m water-depth; South Atlantic, 31˚58’S, 16˚58’E; AMT16 cruise, June, 2005; XP

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